Life on the edge: celebrating a successful long-term ecological study


Photo credit: Ken Wilson

The Scottish isles of St Kilda, off the west coast of the Outer Hebrides, have an important place in my heart. It was on St Kilda where I first realised that not all sheep are boring, where I sustained my first fieldwork-related injury (a broken hip caused by an impact during a sheep-chasing incident!), where I successfully ran my first Research Council grant, and where I met my wife. It was also on St Kilda where I gained my first taste of a long-term ecological study, and where I realised what a tremendous effort is required to ensure their sustained persistence. On my first visit (lambing 1993), the St Kilda Soay sheep project was still in its infancy, having been conceived in its current form by Tim Clutton-Brock and Steve Albon in 1985. This year, the project celebrates its 30th anniversary and to mark this milestone the team organised a programme of public talks at the Royal Society of Edinburgh.

I am sure that a summary of all the excellent presentations at the meeting will appear elsewhere, so here rather than repeat this, I wanted to reflect on some of the broader issues that these talks, and the project as a whole, raises about what makes a successful long-term study (with a special nod to Dave Coltman for the inspiration).

But first, is this really a long-term study and has it really been that successful?

Well, 30 years certainly feels like a long time, but the Soay sheep project is a spring chicken in comparison to some other long-term animal ecology studies – think great tits at Wytham Wood, chimpanzees at Gombe, elephants at Amboseli, Darwin’s finches on Daphne Major, aphids at Rothamsted, red deer on Rum, guillemots on Skomer, to name but a few [see Begging for funding]. However, if success if gauged in terms of research publications then the project compares favourably with the best of these – over the last three decades more than 150 papers have been published from the project, accumulating between them more than 5,000 citations (h-index = 42).

Photo: Ken Wilson

Photo: Ken Wilson

Another useful metric of success is the number of young scientists the project has trained and mentored. This is more difficult to quantify, but at a best guess the project has probably trained several dozen doctoral students and post-doctoral scientists (including two current, and one previous,  JAE senior editors!), not to mention scores of undergraduates and Masters students who have benefited from Soay sheep data and samples, and the literally hundreds of volunteers who have gained a valuable life experience. Of course, the Soay sheep project is not unique in either of these regards – indeed the production of lots of high-quality publications and young scientists is a well characterised output from most long-term studies, which is why they should be promoted and protected, as discussed previously on this blog.

So, why has the Soay sheep project been so successful?

Generous leadership: Over the years, a hallmark of the Soay sheep project has been that it has welcomed in new collaborators who bring in fresh skills and perspectives. What initially started out as an analysis of the causes and consequences of the unstable sheep population dynamics, quickly incorporated new questions about sheep genetics and the inheritance of traits, the role of parasites, vegetation dynamics, sheep behaviour, demography, quantitative genetics, genomics, immunology, ageing and physiology. It would have been easy for the project leaders to guard access to such a valuable and unique ecological resource, but by inviting new scientists to the party and encouraging ex-students and post-docs to develop their own areas for development, the Soay sheep project has continued to be at the vanguard of ecological science, with new collaborators adding value to ongoing studies rather than competing with them. That is not to say that it has always been smooth sailing (either metaphorically or literally – access to the islands requires a sea journey of at least 40 miles on often choppy seas!), or that personalities and egos have not clashed at times over the years. But under the considered and generous leadership, of first Tim Clutton-Brock and then Josephine Pemberton, the long-term future and development of the project has always taken centre stage.

Photo credit: Ken Wilson

Photo credit: Ken Wilson

The long game: Another reason for the project’s continued success is that whilst grant funding is typically short-term (usually three years), the outlook of the project has always been longer-term, with strategic planning of grant applications ensuring continual Research Council funding for the entire 30 years of the project – a quite spectacular feat! This has been made possible only by the combined efforts of lots of individuals and not just the nominal leaders of the project. Science funding is typically fickle, with research ideas coming in and out of fashion. An important bi-product of generous leadership is that there can be a pluralist approach to funding, with multiple applications for core funding being possible due to the diverse nature of the study and the questions that are currently considered ‘sexy’. Indeed, over the years the ‘core’ long-term monitoring of the Soay sheep project has been funded by grants to at least half-a-dozen different individuals.

The project has also taken a long-term approach to its understanding of the key questions being tackled. As ecologists, we are well aware that we can never fully understand ecological systems, and that all we can ever really do is to establish our best guess at the ‘truth’, which we must then update in light of new information. This is perhaps best illustrated by considering how our understanding of Soay sheep population dynamics has evolved since the project began. In 1992, just before I first visited St Kilda, Bryan Grenfell and others used the available data (6 years of high-quality census data from Village Bay) to argue that the unstable dynamics of the sheep were in fact intrinsic cycles driven by overcompensating density-dependent mortality. After I joined the project, we updated that assessment (based on 40 years of whole island census data from Hirta and neighbouring Boreray) to argue that the dynamics were not in fact cyclical but were due to pronounced threshold effects with population crashes occurring in years above some critical sheep density, the depth of which depended on winter weather (as illustrated by synchronous crashes occurring on isolated but neighbouring islands – the Moran effect). A few years later, Tim Coulson and colleagues showed that population age-structure was also important. Oh, and by the way, the long-term trend is for the sheep population size to increase and for the sheep themselves to get smaller, probably due to climate change. And to understand the mechanisms underpinning all this, we also need to consider the interaction between the sheep and their food supply: after more than 20 years of twice-yearly vegetation monitoring by Mick Crawley and students, we are only now getting close to understanding how the sheep and their food supply interact with each other and climate. The point is this: only by combining long-term data collection with a multi-disciplinary research agenda can we hope to tackle these ecosystem-scale interactions and the impact of large-scale phenomena such as global warming.

Photo Credit: Ken Wilson

Photo Credit: Ken Wilson

Continuity: Finally, another factor contributing to the success of the Soay sheep project is the continuity provided not just by its leadership and long-term collaborators, but also by its field staff and data curators. Jill Pilkington MBE has worked with the sheep project for over 20 years now, visiting the island for weeks at a time every spring, summer and autumn, come rain or shine. Ian Stevenson completed his PhD on St Kilda in 1994 and for the past 15 years has developed and managed the Soay sheep database, latterly in his role as MD of Sunadal Data Solutions . Their continued contribution to the project (and in Ian’s case to many other long-term studies) has not only allowed new methods and approaches to be developed and refined to work like clockwork, building on previous successes and rectifying previous errors, but has also provided an invaluable resource for new students and collaborators to mine. It is also worth noting here the important long-term support provided by the National Trust for Scotland, which owns the St Kilda, and Scottish Natural Heritage, which has oversight of conservation on the islands; without their continued understanding of the value of scientific research, this project could have died a long time ago.

I am not sure if the traits I have highlighted above are common to other successful long-term studies or if the Soay sheep project is unique. But either way, I think it provides a valuable example for other potential long-term studies to follow.

Ken Wilson

Senior Editor (@spodoptera007)

Three decades of Ken and Soay sheep!

Three decades of Ken and Soay sheep!

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Penguins on Parade: Conflict in South Georgia – A Slideshow

This gallery contains 16 photos.

The Sub-Antarctic island of South Georgia is home to some of the world’s largest breeding aggregations of penguins. Long-term monitoring studies reveal that the local population trends are complex. Some species and colonies have rapidly declined, but others have increased … Continue reading

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VIDEO In hot and cold water: Life-history biology of the Antarctic Kiwaidae

In 2010, a UK-led expedition to the Southern Ocean revealed a community of deep-sea animals thriving around volcanic vents on the ocean floor near Antarctica. Among the many new species discovered, was the visually abundant yeti crab, Kiwa tyleri. As a result of local thermal conditions at the vents, these crabs are not restricted by the physiological limits that otherwise exclude reptant decapods from the cold stenothermal waters south of the polar front. Using a deep-sea remotely operated vehicle (ROV), research led by the University of Southampton reveals the adult life-history of this species by piecing together variation in microdistribution, body size-frequency, sex ratio, and ovarian and embryonic development, which indicates a pattern in the distribution of female Kiwaidae in relation to their reproductive development. These findings are published in the Journal of Animal Ecology (Marsh et al., 2015, In hot and cold water: differential life-history traits are key to success in contrasting thermal deep-sea environments).

(No audio)

Leigh Marsh
University of Southampton
(twitter: @Leigh_Marsh)

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Chi square I’ve met you before. A belated Valentine’s blog

For those readers who have met me, it will come as no surprise that I was a bit of a geek when I was doing my undergraduate studies.  And that was long before geek was in any way sexy.  Sheldon (from ‘The Big Bang Theory’ and not @ben_sheldon_EGI) probably hadn’t been born.  However, one day one of the cool gang of undergraduates did talk to me.  She wondered whether she could use my results from a practical she had been ‘unable’ to attend.  I wanted to help but I was also concerned she’d copy my data and I’d end up being the one hauled over the coals for plagiarism.  So I came up with a cunning plan.  I wrote some code on the VAX (look it up online if you’re under 45) that took my data and generated a pseudo random dataset with many of the same statistical properties as the dataset I had collected.  It took me most of the night.  Nicole seemed happy, but not sufficiently so to come for a drink with me. Continue reading

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My life as a wealth generation asset

A few years ago, someone with an interest in dynamical systems devised a complex financial product that allowed banks and other larger corporations to achieve a good return on their investments at limited risk. But it turned out that their money was not as safe as they thought, and things went belly up. Banks lost money hand over fist, some ended up bankrupt, while the taxpayer bailed others out. The global economy took a nosedive, and countries ended up being much more in debt than they would have liked. As the next general election approaches, we are told that things are improving in the UK, but the deficit is large, and it is not coming down as quickly as expected. This is a serious problem, and something that will take time to sort out. There must have been many very difficult meetings in Whitehall, with departments told they need to spend less money and, where possible, generate money. Whether one agrees with this strategy or not, the logic behind it is straightforward to follow: we need to pay off our debts so we should spend less money and generate more of it. Continue reading

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How complex should models used by ecologists be?

In his thought-provoking blog, Tim asks a fundamental question every ecologist has to think about occasionally: how many terms should I include in my model? Tim argues that models with a high heuristic value include only a few parameters; models like Verhulst’s logistic model of population dynamics and Lotka-Volterra’s predator-prey model. Tim also advises that ecologists in the quest of universal laws should limit the number of parameters in their models to as few as necessary to get the job done. However, I shall argue that the devil is in the detail! Continue reading

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Modelers to the left of me, field biologists to the right; here I am, stuck in the middle with you

One of the things that I enjoy most about the science I do is collaborating with both field biologists who know their systems inside out, and theoreticians who’s specialist expertise is abstraction and equations. One thing I have learned from these collaborations is that every field or laboratory system exhibits some oddities. The Trinidadian guppy system is the latest, wonderful, system I have begun collaborating on, and it exhibits numerous quirks. One of my favourites is what we affectionately term ‘zombie males’. Because females store sperm, males can sire offspring after death. Such behavior is, of course, not particularly unusual, but this is the first time I have had to ponder whether it is necessary to incorporate such a life history ‘quirk’ in models, and if so, how. These system-specific oddities make me take issue with a quote from a theoretician colleague. It goes something like this: ‘reality is just a special case, and not a particularly interesting one’. Reality is, in fact, very interesting.   However, the oddities of each system do generate certain challenges for the modeler. Should they always be incorporated into models? Continue reading

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Down the up staircase: longevity and academics

Forty years. That’s at least how long an academic career can last, if you start at 30 and retire at 70. There is no mandatory retirement age (at least in the UK and US) and, unlike most people, tenured academics rarely lose their jobs. For older academics (say over 50) increased longevity can be accompanied by the right to work as long as one wants.

The usual career pattern – always sideways or up, rarely down – means that academics spend 20 years at near-maximum salary and with a tight grip on institutional power and hiring practices. This isn’t actually bad for productivity since studies convincingly show that aging doesn’t affect productivity. And the ability to work into old age is attractive to researchers whose salaries often lag the business sector. Continue reading

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Biodiversity v Intensive Farming; Has Farming Lost its Way?

Modern intensive farming produces plentiful, cheap food but is reliant on heavy use of agrochemicals and is a major driver of the ongoing collapse of wildlife populations. Taxpayers pay billions each year to support this system, with the bulk of this money going to the biggest, richest farming operations. In this blog I examine how we got to this unhappy position, question the need to further increase food production given current food waste, and suggest that we need to move towards a more sustainable, evidence-based farming system, with a source of independent advice for farmers, rather than allowing the agrochemical industry to shape the future of farming.

It is not politically correct to criticise farmers or farming. We are brought up on stories about the adventures of a playful piglet who lives on a farm with a sheepdog, half a dozen chickens and a smiling cow, all presided over by a rosy-cheeked farmer, his wife and their two children. Farmers might also be portrayed as custodians of the land, where the countryside that they look after is filled with the sound of skylarks singing, bumblebees buzzing amongst the hedgerows, and butterflies flitting across sunlit, flowery meadows.

Farming is of course the most fundamentally important of human activities; without farms and farmers, we would quickly starve. Going back to hunter-gathering is not an option. What is more, the human population is growing, and therefore we must increase food production. The United Nations Food and Agriculture Organisation (FAO) declared in 2010 that we must double food production by 2050, and this rationale is used to justify the drive for ever-increasing yield. One might argue that we should focus all our research on increasing yield at all cost, else our grandchildren will starve. Continue reading

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A Look Back At 2014

It’s been a busy old year at Journal of Animal Ecology, with lots of personnel changes and a few new initiatives. Here, we review some of these developments.

The blog

In terms of new initiatives, the highest profile is arguably this blog – Animal Ecology In Focus – which we started in June 2014. Although the senior editors were initially quite sceptical about whether this latest venture into social media would be successful, the feedback we have had so far suggests that it is a valued addition to our outputs, along with our Twitter feed (@AnimalEcology), our Facebook page, podcasts, videos, etc. The blog was kick-started by a controversial post by our own Tim Coulson on the latest UK badger cull trials and this theme was picked up again in a later post by the other Senior Editors, who offered the services of the Journal to Defra to provide an independent assessment of this year’s badger cull trial. The blog was subsequently highlighted by the BBC and cited in a Westminster debate by MPs from across the political divide. It is likely that this issue will continue to feature on our blog for some time to come. Other notable posts in the last six months include one by Ken Wilson highlighting the decline in entomological papers published in the Journal over the last 40 years, one by Ben Sheldon on funding long-term studies and a number of posts by Tim Coulson on issues such as sex-biases in science, the value of archiving data (with Ben), and a call for pre-proposals. In December, we opened up the blog for the first time to our Associate Editors, with a powerful post by Sonia Altizer and Julie Rushmore on the role of wildlife in the spread of Ebola virus. In 2015, we plan to invite other renowned experts in animal ecology to share their thoughts with on a range of topical issues. The first of these, by Dave Goulson, will appear shortly. If you have any ideas about what might make an interesting blog post, and who might write it, please contact us. Continue reading

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The Wildlife Side of Ebola: What Animal Ecology Can Contribute to Studying the Spread of a Deadly Virus

chimpanzee troop 6 v2

Chimpanzees in Kibale National Park, Uganda. Photo: Julie Rushmore.

Ebola virus as a zoonotic pathogen

The current Ebola virus outbreak in West Africa is very much on people’s minds as a story of human suffering and death, with nearly 15,000 Ebola cases reported from West Africa as of November. Ebola virus spreads rather slowly but causes a remarkably high fatality rate, with 50% or more of human cases ending in death. The current epidemic dwarfs the cumulative number of cases in previous localized outbreaks, motivating new research into vaccines, treatments, and efforts to slow Ebola spread. Promising signs of slowing transmission have emerged in recent weeks, but many experts predict that widespread vaccination will be needed to fully halt the epidemic. The social, political and economic impacts of the current Ebola virus epidemic will likely linger for years to come. Continue reading

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It is alright to be wrong and was Wright right?

Max Planck famously said ‘science advances one funeral at a time’. Sadly there is still some truth to this: some scientists are incapable or unprepared to change their views despite overwhelming scientific evidence that they are wrong*. Outdated ideas often only die with their advocates. One thing I try to teach students is that it is alright to be wrong: many ideas turn out to be incorrect, lots of exciting hypotheses end up not being supported, and frequent promising avenues turn out to be cul-de-sacs. But that is how science progresses. We need to rule out some competing hypotheses in order to advance knowledge. Continue reading

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Archive your data!

When you submit a manuscript to the Journal of Animal Ecology you are asked the following question: ‘If your paper is accepted for publication where do you expect to archive your data or, if already archived where are the data held?’ Recently, we received a manuscript where the author had responded to this question with: ‘the data will be archived with the lead author’. This is not an appropriate digital data archive!

On about the same date that the previous was submitted, one of us wrote to the authors of a paper published just over a decade ago. In the email, data underpinning the paper were requested, as it seemed plausible that the paper’s conclusions were a consequence of an error in analysis. The authors were unable to provide the data because they had changed computers several times in the last decade and it had been lost somewhere along the line. This won’t be the only time this has happened. In fact, not so long ago, one of us has had to play the embarrassing role of replying to request a for data with the news that it was in a file format that was inaccessible on any current operating system. Fortunately, these sort of things shouldn’t happen in future for Journal of Animal Ecology papers as one of the reasons we require authors to upload data associated with their papers to a respected data repository is to avoid just such scenarios. Continue reading

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A Call for Pre-Proposals

This year I have written two UK research council proposals and a European Research Council grant. They are each on completely different topics. I suspect it has taken about six months of my time. I was pleased with each application, but I don’t have high hopes for any of them, simply because funding rates are low. I am not atypical, and this is not an atypical year for me.

What happens at the UK research councils is one receives reviewer comments on the proposal. From my experience, about half of the reviews are positive suggesting funding, and the remaining ones grumble. They rarely raise any scientific objectives that cannot very easily be dealt with. They often criticize the research team, complain that some key literature is missing and request additional methodological details. A colleague of mine once told me that he had had several grants rejected at NERC that were better than everything he had ever been asked to review, and consequently always wrote grumpy reviews. He is a little delusional. Anyway, once reviewer comments are received, a response can be written. The committee assesses all grants, reviewer comments and responses, ranks proposals and the top few get funded. Continue reading

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Begging for funding?

Understanding ecological systems takes time. While some experimental ecological work, performed under controlled lab conditions, can be conveniently fitted into the short-term periods beloved of funding bodies, much of ecology requires a longer-term perspective. Why is that? First, life-histories frequently operate at generational scales approaching decades. To have any hope to make sense of patterns of inheritance, selection or demography we need data spanning multiple generations, and that may mean multiple decades. Second, almost all ecological studies reveal heterogeneity among individuals – frequently in terms of vital rates, or detection probability, or other aspects of life-histories. Such heterogeneity makes it very hard to extrapolate from cross-sectional observations to understand the true sources of variation driving a population. Continue reading

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Transparency and Evidence-Based Policy: An Open Letter to Defra from Journal of Animal Ecology

As a scientific journal, we are in the business of independently assessing the rigour of work conducted by the research community, including the methods it uses to collect, analyse and interpret appropriate data. We are therefore well placed to judge the merits of relevant scientific endeavour and to provide constructive feedback. On October 30th 2014, the UK’s Shadow Farming Minister, Huw Irranca-Davies, called for an independent review of the methods being used to assess the outcomes of the ongoing pilot badger culls in England 1. Such a review requires a detailed understanding of the behaviour, dynamics and management of wild animal populations – disciplines that are at the heart of the field of animal ecology. As the UK’s leading animal ecology journal, we hereby offer our services to the Secretary of State to provide an independent assessment of the methods and data collected as part of this year’s badger cull. Continue reading

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Solving the skewed sex ratio problem in science

In 2003 Milner-Gulland et al. wrote a paper on extreme adult sex ratios in saiga antelope. Males had become so rare in some years that the behavior of the system became dysfunctional and population performance suffered catastrophically. The only other environments where I know of heavily skewed adult sex ratios are university science faculties. Except here the skew is in the other direction, with females being rare. Social scientists have shown that skewed sex ratios in the workplace can negatively impact many performance metrics (e.g. Fenwick and Neal 2001).

Many scientists are rightly concerned by the paucity of women on the faculty of many science departments, and there has been much contemplation on the causes of attrition as more men progress from Ph.D. to post-doc to a faculty position to full professor than women. There are hypotheses proposed to explain this ranging from men being more likely than women to express the traits thought to aid success in academia including self-belief and an ability to brush off criticism, through to a lack of adequate home life provision. However, identification of these causes does not seem to be having much of an effect on reducing the skewed sex ratio. For example, of 43 researchers offered prestigious Royal Society University Research Fellowships this year, 41 were men (see here). I am not entirely surprised by this. Many ‘solutions’ I have heard proposed to address the skewed sex ratio problem seem unlikely to succeed. For example, one popular call is for women’s groups to be set up. No one has ever succeeded in explaining to me how that is supposed to lead to change. Continue reading

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Where have all the insects gone?

Recently, we commissioned one of Journal of Animal Ecology‘s most experienced Associate Editors, Simon Leather, to compile a Virtual Issue on his great passion – insects. The journal has published many classic insect ecology papers over the years and Simon does a great job of highlighting some of these as well as many new papers that we hope will go on to become classics themselves. In his preface to the VI, Simon bemoans the fact that back in the 1970s, when he first began subscribing to the journal, there were many more papers on insects than there are now and that the journal has perhaps become vertebrate-centric in recent years.

This got me thinking – is this really true? And if it is, then why do we publish fewer entomological papers now than back then? Are we alone in this trend or is it common across other general ecological journals? And, either way, should we be worried about the taxonomic distribution of our papers? Continue reading

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Name: UK government. Animal ecology test score: 0

© Andrew Byrne

© Andrew Byrne

Every now and again animal ecology findings make it into the news. Press coverage often focuses on cases where a species is on the edge of extinction, has erupted to plague proportions, or exhibits some quirky behaviour. One of the positive things about such coverage is that the public appreciates that animal ecology is a mature field of study that uses high-tech methods of data collection, cutting-edge statistical methods and mathematically elegant models. But all too often animal ecology stories are little more than a curiosity, chosen to fill the ‘And finally…’ slot. Occasionally animal ecology research influences government policy – something that has happened with the control of tuberculosis (TB) in cattle. However, this particular case is not a good news story – sound animal ecology advice is being ignored by the current UK government. The reason? A cynic might speculate that it is because following best animal ecology practice might lead to conclusions at odds with what the government seems unjustifiably determined to do. Continue reading

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